Morphological phylogeny of cobweb spiders and their relatives (Araneae, Araneoidea, Theridiidae) Author Agnarsson, Ingi text Zoological Journal of the Linnean Society 2004 2004-08-31 141 4 447 626 https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2004.00120.x journal article 10.1111/j.1096-3642.2004.00120.x 0024-4082 5431156 PLACEMENT OF THERIDIIDAE AND ITS RELATIVES This analysis tested the placement of Synotaxus (formerly an argyrodine theridiid) in a matrix containing argyrodine theridiid genera. In concordance with Forster et al. (1990) , Griswold et al. (1998) and Agnarsson (2003c) , Synotaxus is not a theridiid. The web, egg-sac guarding web and general somatic morphology make Synotaxus unlike any theridiid ( Fig. 95A–D ). Instead, it is sister to the theridioids (clade 55), as also suggested by Griswold et al. (1998) , who informally named the clade containing cyatholipoids (cyatholipids plus synotaxids) and theridioids as the spineless femur clade (clade 57). Forster et al. (1990) expressed doubts concerning the position of Carniella within Theridiidae , based on the basal position of the ‘paracymbium’ ( Fig. 36C ) and the lack of flattened AG spigots ( Fig. 36I ) so typical of theridiids. However, the results of this study strongly support such a placement (see Thaler & Steinberger, 1988 ) as sister to the clade ( Pholcomma , Robertus ) ( Fig. 102 ). Knoflach (1996) suggested affinities between Carniella and Theonoe , based on the absence of the male palpal tibial trichobothrium and modification of the cymbial tip ( Fig. 36A–C ). Theonoe shares several additional features with the clade containing Carniella , Pholcomma and Robertus and thus Knoflach’s argument seems well founded. Table 2. Species autapomorphies, numbers refer to characters; synapomorphic characters are mapped in Figures 103 and 104

Species	Autapomorphic characters
Argiope argentata	111, 112, 170
Tetragnatha extensa	10, 14, 30, 71, 102, 113, 119, 144, 145, 209, 226, 228, 238, 241
Pimoa rupicola	21, 30, 41, 108, 111, 122, 178, 191, 200, 211, 222
Linyphia triangularis	1, 8, 11, 19, 30, 62, 71, 97, 104, 113, 119, 135, 144, 186, 189, 214, 223, 237, 239, 242
Synotaxus monoceros	131
Synotaxus waiwai	71
Eidmanella pallida	62, 163
Dipoena nigra	4, 45, 201
Euryopis gertschi	19, 174
Thwaitesia margaritifera	8, 42, 51, 65, 146, 212, 225, 226, 233
Episinus amoenus	5, 7, 59, 108
Spintharus flavidus	33, 38, 78, 104, 108, 168, 179
Latrodectus geometricus	231
Crustulina sticta	18, 19, 24, 90, 135, 140, 161, 178, 179, 187
Steatoda grossa	5, 169, 189, 223, 240
Carniella siam	5, 32, 80, 89, 111, 131, 132, 168, 185, 190, 215, 222
Robertus frontatus	8, 11, 83, 89, 98, 111, 152, 156, 219
Pholcomma hirsutum	56, 58, 72, 123, 166, 222
Enoplognatha ovata	135
Selkirkiella magallanes	143
Phoroncidia sp.	16, 17, 28, 57, 58, 78, 130, 135, 137, 142, 143, 149, 150, 167, 172, 176, 182, 225
Cerocida strigosa	11, 51, 108, 112, 121, 123, 150, 163, 168, 173, 190, 210
Stemmops nr. servus	19, 25, 35, 62, 83, 84, 87, 89, 91, 184, 200
Neospintharus trigonus	64, 90, 135, 142
Rhomphaea metaltissima	15, 125, 168, 195, 231
Ariamnes attenuatus	7, 11, 24, 96, 128, 138, 231
Faiditus cf. chickeringi	6, 7, 85, 142, 185
Kochiura aulica	85
Kochiura rosea	81, 111
Anelosimus analyticus	188
Anelosimus cf. jucundus	46, 170
Anelosimus eximius	99
Anelosimus vittatus	156
Chrysso cf. nigriceps	6, 38, 70, 76, 128, 148, 151, 178, 201, 205
Nesticodes rufipes	7, 161, 177, 189
Theridion longipedatum	110, 111, 140
Theridion varians	146, 160
Coleosoma floridanum	41, 76, 135, 152
Tidarren sisyphoides	23, 33, 35, 45, 51, 78, 86, 129, 147, 151, 152, 226, 236
Helvibis cf. longicaudatus	7, 8, 41, 55, 63, 66, 69, 91, 123, 152, 199, 201
Thymoites unimaculatum	1, 137, 143, 178, 190
Ameridion sp.	115
Theridula emertoni	33, 56, 62, 80, 97, 125, 152, 178, 218, 223
Achaearanea tepidariorum	94, 108, 240

As suggested by Berland (1932) , Gertsch (1949) , Wunderlich (1978) and Forster et al. (1990) hadrotarsines are related to theridiids, and here form the sister clade to the remaining theridiids. Hadrotarsine monophyly is supported by an array of morphological peculiarities, making them unlike any other theridiid. They also lack several characteristics typical of other theridiids (see below). For these reasons, revalidating Hadrotarsidae seems logical. However, such an act has major nomenclatural consequences and is currently contradicted by molecular evidence (see Arnedo et al. , 2004 ). Therefore, the elevation of Hadrotarsinae to family rank will be reconsidered following combined analysis (see ‘Note’ above).